Evidence for competition between carnivorous plants and spiders

Several studies have demonstrated that competition between disparate taxa can be important in determining community structure, yet surprisingly, to our knowledge, no quantitative studies have been conducted on competition between carnivorous plants and animals. To examine potential competition between these taxa, we studied dietary and microhabitat overlap between pink sundews (Drosera capillaris) and wolf spiders (Lycosidae) in the field, and conducted a laboratory experiment examining the effects of wolf spiders on sundew fitness. In the field, we found that sundews and spiders had a high dietary overlap with each other and with the available arthropod prey. Associations between sundews and spiders depended on spatial scale: both sundews and spiders were found more frequently in quadrats with more abundant prey, but within quadrats, spiders constructed larger webs and located them further away from sundews as the total sundew trapping area increased, presumably to reduce competition. Spiders also constructed larger webs when fewer prey were available. In the laboratory, our experiment revealed that spiders can significantly reduce sundew fitness. Our findings suggest that members of the plant and animal kingdoms can and do compete.     

Prey‐specific foraging tactics in a web‐building spider

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Various competitive interactions explain niche separation in crop‐dwelling web spiders

Competition for resources is a major organizing principle in communities of organisms that share similar ecological niches. Niche separation by means of exploitation or interference competition was investigated in two taxa of crop‐inhabiting spiders that overlap in microhabitat use and have similar web design. Competition for prey and web sites was tested in microcosm experiments with the most common species that build sheet‐webs: Enoplognatha gemina (Theridiidae) and Alioranus pastoralis (Linyphiidae). A field survey over the crop season provided data on spatial and temporal dispersion of Enoplognatha spp. (Theridiidae) and linyphiid spiders (Linyphiidae) and on availability of prey over the season. In the microcosm experiments, both taxa took springtails as prey, but only Enoplognatha fed on aphids. Differences in diet, however, could not be attributed to either exploitative or interference competition. Spatial separation of websites was attained by vertical displacement of webs in the vegetation (Enoplognatha) and by avoidance of patches occupied by conspecific or heterospecific individuals (linyphiids). In the field, densities of linyphiids and Enoplognatha were correlated negatively and webs were over‐dispersed relative to a random distribution. Both taxa colonized the field at the start of the season; linyphiids colonized as adults, quickly reproduced, and had a second adult peak; Enoplognatha matured in the middle of the season and their numbers remained fairly constant over the season. The combined experimental manipulations and field data suggest that niche separation occurs at different scales. The hypothesis of competition for websites was partially supported, while prey preference (or tolerance) and temporal differences in life history stages also may explain the negative correlations between densities of the two taxa.     

Distribution of plant‐dwelling spiders: Inflorescences versus vegetative branches

Abstract The influence of the architecture of vegetative branches on the distribution of plant‐dwelling spiders has been intensively studied, and the effects on the aggregation of individuals in several spider species on plants include variation in prey abundance, availability of predator‐free refuges and smoother microclimate conditions. The emergence of inflorescences at the reproductive time of the plants changes branch architecture, and could provide higher prey abundance for the spiders. The distribution of spiders between inflorescences and vegetative branches was compared on four widespread plant species in a Brazilian savannah‐like system. Inflorescences attracted more spiders than vegetative branches for all plant species sampled. The influence of branch type (inflorescence and vegetative) on spider distribution was also evaluated by monitoring branches of Baccharis dracunculifolia DC. in vegetative and flowering periods for 1 year, and through a field experiment carried out during the same period where artificial inflorescences were available for spider colonization. Artificial inflorescences attached to B. dracunculifolia branches attracted more spiders than non‐manipulated vegetative branches for most of the year. However, this pattern differed among spider guilds. Foliage‐runners and stalkers occurred preferentially on artificial inflorescences relative to control branches. The frequencies of ambushers and web‐builders were not significantly different between treatment and control branches. However, most ambush spiders (65%) occurred only during the flowering period of B. dracunculifolia, suggesting that this guild was influenced only by natural inflorescences. The experimental treatment also influenced the size distribution of spiders: larger spiders were more abundant on artificial inflorescences than on vegetative branches. The hypothesis that habitat architecture can influence the spider assemblage was supported. In addition, our observational and experimental data strongly suggest that inflorescences can be a higher quality microhabitat than non‐reproductive branches for most plant‐dwelling spiders.     

Choosing Hunting Sites: Web Site Preferences of the Orb Weaver Spider,Neoscona crucifera,Relative to Light Cues

Field experiments carried out on the nocturnal orb weaver spider, Neoscona crucifera (Aranea: Araneidae), found in deciduous hardwood forests suggest that lighted areas where prey densities are elevated provide cues used by the spiders to rank optimal foraging sites. Specifically, experiments were conducted to test whether spiders exhibited preferences for lighted areas where prey densities are high, maximizing their energy intake per unit of foraging time, and minimizing energy expended on web building. Incandescent light bulbs of 4–60 W were used to influence prey densities, and results indicate that when given a choice of brighter versus darker foraging areas, spiders seek lighted areas where prey densities are high. In addition, results support the hypothesis that the size and time of web construction are drastically reduced in brighter situations.     

Different groups of ground-dwelling spiders share similar trophic niches in temperate forests

1. Generalistic interactions between predator and prey may vary with ecosystem type, predator traits, and prey traits, but the interplay of these factors has not been assessed in ground food webs.
2. We investigated trophic interactions of ground-dwelling spiders across eight forests in European Russia associated with body size, hunting strategy, microhabitat specialization, potential prey type, potential prey population density, and forest type (coniferous vs. broadleaved). We analyzed 128 individual spiders, including juveniles, all identified to the family level with two complementary methods: molecular gut content analysis, and stable isotope analysis of carbon and nitrogen.
3. The results suggest that feeding frequency of spiders is affected by predator body size and by selection of certain prey type. Stable isotope analysis showed similar trophic niches among spider families, varying moderately with forest type. Larger spiders had higher Δ¹³C values than smaller ones, but similar Δ¹⁵N values, suggesting that different size classes of spiders belong to different food chains. Results based on stable isotope and molecular gut content analyses were weakly linked, indicating them targeting different trophic niche dimensions.
4. At least for the group-level interactions, family identity and hunting strategy of predator has little predictive power while predator body size and prey traits affected trophic niche dimensions calling for future studies in this direction. Large spiders feed more and rely on different basal resources than small spiders, suggesting that including small species and juveniles provides a more comprehensive picture of food web organization.

     

Pennycress double‐cropping does not negatively impact spider diversity

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Competition for a limited space in kleptoparasitic Argyrodes spiders revealed by field experiments

Most kleptoparasitic Argyrodes spiders rely exclusively on host spider webs for obtaining their food. Because their densities occasionally reach high levels within a restricted area, competitive interactions may be important for determining the density of these unique spiders. Here I used two Argyrodes species commonly found on webs of the large orb-web spider Nephila clavata to clarify whether inter- and intraspecific competition influences abundance and within-web distribution by using observational data and field experiment. Removing Argyrodes flavescens from the host webs induced a remarkably high immigration of that species while density on control webs remained almost at the same level, which is evidence for strong intraspecific competition. Larger individuals of A. flavescens were located more frequently at the capture area of the host webs where it is easy to access prey ignored by the host spider, and spiders immigrating into webs from which that species had been removed were smaller in body size, suggesting interference competition for space among conspecific kleptoparasites. Argyrodes bonadea increased in number on webs from which A. flavescens had been removed, and the increase was correlated with the number of A. flavescens removed. This finding is evidence for interspecific competition that is rarely reported in spiders. A multiple regression model including numbers of a conspecific parasite as well as web and body sizes of the host spider could not detect competitive interactions between species, suggesting the importance of experimental approaches. Received: May 22, 2000 / Accepted: December 1, 2000     

Guiding lights: Foraging responses of juvenile nocturnal orb‐web spiders to the presence of artificial light at night

The reach of artificial light at night (ALAN) is growing rapidly around the globe, including the increasing use of energy‐efficient LED lights. Many studies document the physiological costs of light at night, but far fewer have focused on the potential benefits for nocturnal insectivores and the likely ecological consequences of shifts in predator–prey relationships. We investigated the effects of ALAN on the foraging behaviour and prey capture success in juvenile Australian garden orb‐web spiders (Eriophora biapicata). Laboratory experiments demonstrated that juvenile spiders were attracted to LED lights when choosing foraging sites, but prey availability was a stronger cue for remaining in a foraging site. Field experiments revealed a significant increase in prey capture rates for webs placed near LED lights. This suggests that any physiological costs of light at night may be offset by the foraging benefits, perhaps partially explaining recently observed increases in the size, fecundity and abundance of some orb‐web spider species in urban environments. Our results highlight the potential long‐term consequences of night lighting in urban ecosystems, through the impact of orb‐web spiders on insect populations.     

Does the strength of cross‐ecosystem trophic cascades vary with ecosystem size? A test using a natural microcosm

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Niches and interspecific competitions of wolf spiders and mirid bug, Cyrtorhinus lividipennis under three pest management strategies in paddy fields

The duck-rice and fish-rice were two important strategies of pest management in the process of organic rice. The niches and interspecific competitions of wolf spiders and mirid bug, Cyrtorhinus lividipennis under three strategies of pest management in paddy fields were studied in order to understand the influences of different pest control methods on ecological relationships between the two main types of predators. The results of our study suggested that ducks might prey upon wolf spiders resting on the base of rice plants, thus the populations of wolf spiders decreased in duck-rice treatment. The impact of ducks on mirid bug was not significant because it preferred to congregate on the mid-part of rice plants. The temporal selectivity of wolf spiders was smaller than that of mirid bug, which meant occurrence frequency of wolf spiders was higher compared with mirid bug. In addition, competition between wolf spiders and mirid bug was significantly more intense in duck-rice or fish-rice treatment than in natural control treatment at different stages. Probably, ducks and fishes drove wolf spiders away from their original spatial niche, the base to the mid-part in rice plants, enhancing competition between wolf spiders and mirid bug in duck-rice or fish-rice treatment. Based on the results of this study, duck-rice or fish-rice, as a control pest method in the process of organic rice, should be prudentially treated.     

Caught in the web: Spider web architecture affects prey specialization and spider–prey stoichiometric relationships

Quantitative approaches to predator–prey interactions are central to understanding the structure of food webs and their dynamics. Different predatory strategies may influence the occurrence and strength of trophic interactions likely affecting the rates and magnitudes of energy and nutrient transfer between trophic levels and stoichiometry of predator–prey interactions. Here, we used spider–prey interactions as a model system to investigate whether different spider web architectures—orb, tangle, and sheet‐tangle—affect the composition and diet breadth of spiders and whether these, in turn, influence stoichiometric relationships between spiders and their prey. Our results showed that web architecture partially affects the richness and composition of the prey captured by spiders. Tangle‐web spiders were specialists, capturing a restricted subset of the prey community (primarily Diptera), whereas orb and sheet‐tangle web spiders were generalists, capturing a broader range of prey types. We also observed elemental imbalances between spiders and their prey. In general, spiders had higher requirements for both nitrogen (N) and phosphorus (P) than those provided by their prey even after accounting for prey biomass. Larger P imbalances for tangle‐web spiders than for orb and sheet‐tangle web spiders suggest that trophic specialization may impose strong elemental constraints for these predators unless they display behavioral or physiological mechanisms to cope with nutrient limitation. Our findings suggest that integrating quantitative analysis of species interactions with elemental stoichiometry can help to better understand the occurrence of stoichiometric imbalances in predator–prey interactions.     

Parasitoid genus‐specific manipulation of orb‐web host spiders (Araneae,Araneidae)

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Intraguild interactions between spiders and ants and top-down control in a grassland food web

In most terrestrial ecosystems ants (Formicidae) as eusocial insects and spiders (Araneida) as solitary trappers and hunters are key predators. To study the role of predation by these generalist predators in a dry grassland, we manipulated densities of ants and spiders (natural and low density) in a two-factorial field experiment using fenced plots. The experiment revealed strong intraguild interactions between ants and spiders. Higher densities of ants negatively affected the abundance and biomass of web-building spiders. The density of Linyphiidae was threefold higher in plots without ant colonies. The abundance of Formica cunicularia workers was significantly higher in spider-removal plots. Also, population size of springtails (Collembola) was negatively affected by the presence of wandering spiders. Ants reduced the density of Lepidoptera larvae. In contrast, the abundance of coccids (Ortheziidae) was positively correlated with densities of ants. To gain a better understanding of the position of spiders, ants and other dominant invertebrate groups in the studied food web and important trophic links, we used a stable isotope analysis (¹⁵N and ¹³C). Adult wandering spiders were more enriched in ¹⁵N relative to ¹⁴N than juveniles, indicating a shift to predatory prey groups. Juvenile wandering and web-building spiders showed δ¹⁵N ratios just one trophic level above those of Collembola, and they had similar δ¹³C values, indicating that Collembola are an important prey group for ground living spiders. The effects of spiders demonstrated in the field experiment support this result. We conclude that the food resource of spiders in our study system is largely based on the detrital food web and that their effects on herbivores are weak. The effects of ants are not clear-cut and include predation as well as mutualism with herbivores. Within this diverse predator guild, intraguild interactions are important structuring forces.     

Generalist predators in organically and conventionally managed grass‐clover fields: implications for conservation biological control

Organically managed agroecosystems rely in part on biological control to prevent pest outbreaks. Generalist predators (Araneae, Carabidae and Staphylinidae) are a major component of the natural enemy community in agroecosystems. We assessed the seasonal dynamics of major generalist predator groups in conventionally and organically managed grass–clover fields that primarily differed by fertilisation strategy. We further established an experiment, manipulating the abundant wolf spider genus Pardosa, to identify the importance of these predators for herbivore suppression in the same system and growth period. Organic management significantly enhanced ground‐active spider numbers early and late in the growing season, with potentially positive effects of plant cover and non‐pest decomposer prey. However, enhancing spider numbers in the field experiment did not improve biological control in organically managed grass–clover fields. Similar to the survey results, reduced densities of Pardosa had no short‐term effect on any prey taxa; however, spider guild structure changed in response to Pardosa manipulation. In the presence of fewer Pardosa, other ground‐running spiders were more abundant; therefore, their impact on herbivore numbers may have been elevated, possibly cancelling increases in herbivore numbers because of reduced predation by Pardosa. Our results indicate positive effects of organic farming on spider activity density; however, our survey data and the predator manipulation experiment failed to find evidence that ground‐running spiders reduced herbivore numbers. We therefore suggest that a positive impact of organic fertilisers on wolf spiders in grass–clover agroecosystems may not necessarily improve biological control when compared with conventional farming.     

Contrasting effects of habitat fragmentation,population density,and prey availability on body condition of two orb‐weaving spiders

1. Habitat fragmentation is a major threat to biodiversity because it disrupts movement between habitat patches. In addition, arthropod fitness may be reduced in fragmented habitats, e.g. due to reduced prey availability. 2. We studied the relationship of spider body condition with habitat fragmentation, population density, and prey availability. We expected that prey availability and population density of spiders would be affected by landscape composition and patch isolation. Body condition should be enhanced by high prey availability, but negatively affected by population density due to competition. 3. We sampled spiders on 30 groups of cherry trees that varied independently in the level of isolation from other woody habitats and in the percentage of woody habitat within 500 m radius. As a measure of body condition, we used residuals of the relationship between individual body mass/opisthosoma width and prosoma width of the two most common orb‐weaving spider species, Nuctenea umbratica Clerck and Araniella opisthographa Kulczynski. 4. Body condition of A. opisthographa was positively correlated with the abundance of flies, which increased with the percentage of forest in the landscape. In contrast, body condition of N. umbratica was reduced at high population densities, presumably due to intraspecific competition. In addition, body condition and population density of A. opisthographa was lower at isolated sites. 5. Our study suggests that effects of landscape fragmentation on body condition vary strongly between spider species, depending on the relative role of food limitation and intraspecific competition.     

Presampling sensory information and prey density assessment by wolf spiders (Araneae, Lycosidae)

Decisions regarding foraging patch residence time and the assessmentof patch quality may be mediated by various sources of information.This study examined the use of sensory cues by hunting spidersto assess prey density in the absence of prey capture. Adultfemale wolf spiders [Schizocosa ocreata (Hentz); Lycosidae]had food withheld for 4 days and then were exposed to artificialforaging patches containing four densities of crickets (0, 3,10, 20) with different sensory stimuli (visual and vibratoryinformation, visual only, and vibratory only). The spiders werenot allowed to feed during trials, and patch residence timewas recorded. The spiders varied patch residence time basedon sensory cues alone and spent more time in patches with higherprey density. With visual information only, spiders could apparentlydistinguish among prey densities almost as well as with visualand vibratory cues combined, but residence time did not differamong prey densities when only vibratory information was presented.Measurements of vibration levels produced by cricket activityunder experimental sensory treatments conform to test results,suggesting that visual detection of crickets is important inpatch assessment used in determining patch residence time.     

Habitat selection in a large orb‐weaving spider: vegetational complexity determines site selection and distribution

1. The distribution of the large orb‐weaving spider Argiope trifasciata in old field habitats of North America and the habitat selection process this species used was studied for 2 years. 2. Because web spiders have limited dispersal abilities and an energetically costly prey capture device, they do not have the ability to sample potential foraging sites. Structural complexity of the vegetation to which the web must be attached is relatively easy to assess. The hypothesis that the structural complexity is a primary factor in determining initial web site selection was tested both by relating the natural distribution of the spiders across habitats to vegetational complexity and by manipulating the complexity of the habitats in a series of experiments. 3. Argiope trifasciata was not distributed evenly among three old field vegetation types. Habitat complexity was related to spider density in both years although no measure of insect activity, prey capture, or prey consumption was correlated with spider distribution. 4. Three experimental manipulations were conducted to test the impact of habitat structure on spider establishment: (1) the amount of natural vegetation was reduced, (2) structures were added to a simple habitat, and (3) the complexity of the structures added was varied. In each case, spiders were introduced and establishment of webs was monitored. In all manipulations, spider establishment was related to the complexity of the substrate available. 5. These results are important for understanding the cues that influence foraging site selection and therefore provide insight into the distribution of species with limited dispersal abilities and high site investment requirements.     

When carnivores collide: a review of studies exploring the competitive interactions between bobcats Lynx rufus and coyotes Canis latrans

1. Bobcats Lynx rufus and coyotes Canis latrans are two widespread mesopredators with a complex history of sympatry. The competitive interactions between these species are of interest to biologists due to the furbearer status of bobcats, the recent range expansion of coyotes, and the recolonisation of several parts of North America by bobcats following their extirpation. Although studies exploring the dynamics and competition between bobcats and coyotes span decades, there is a lack of understanding regarding what factors influence exploitative or interference competition, and what methodologies are conducive to identifying these types of competition.
2. We gathered a comprehensive list of research papers (n = 41) exploring bobcat–coyote competitive interactions in North America. From them, we collected the following: study site characteristics, number and types of research methods, number and types of metrics explored, history of sympatry of the two species at the study location, presence of apex predators, and documentation of interference and/or exploitative competition.
3. Using generalised linear models, we determined that interference competition between bobcats and coyotes was observed primarily in open habitat. However, habitat heterogeneity, the number of research methods and metrics used, presence of an apex predator, and history of sympatry could not be used to predict the occurrence of interference competition. Studies that included diet overlap were less likely to observe interference competition than studies that used other metrics to infer competition.
4. Competitive interactions between coyotes and bobcats are largely a function of prey availability. Our findings suggest that habitat type may be a surrogate for prey availability, which many researchers mention, but do not explicitly measure. Future studies investigating bobcat–coyote interactions should include the quantification of prey densities to gain a more comprehensive understanding of the system at large, and should avoid using solely diet or habitat overlap as metrics to assess competition.

     

Functional responses of three guilds of spiders: Comparing single‐ and multiprey approaches

Spiders are successful natural enemies of pests occurring throughout the different strata of an agroecosystem. The study of their functional responses can provide information related to the potential effectiveness of different species and guilds on reducing a pest population. However, multiple prey availability may change the functional response of a predator. In this study, the functional responses of three species of spiders in single‐prey and multiple‐prey experiments were modelled. The spider species Haplodrassus rufipes, Araniella cucurbitina and Synema globosum were chosen as being representatives of ground runners, orb‐weavers and ambushers, respectively. Three prey species were selected: a target prey, the Mediterranean fruit fly Ceratitis capitata and two alternate prey species, the flour moth Ephestia kuehniella and the house cricket Acheta domesticus. When the selected target prey C. capitata was supplied in the single‐prey experiments, the most and the least efficient spider species were H. rufipes and S. globosum, respectively. However, opposite results were obtained when alternative preys were supplied. Also, A. cucurbitina significantly changed its functional response with the presence of alternative prey species. Considering the prey preference during the multiple‐prey experiments, A. domesticus, used as representative of a heavy, long‐sized and highly motile prey, was avoided whereas C. capitata, the target pest used as representative of light, small‐sized and moderately motile prey was preferred by the three species of spiders. Ephestia kuehniella, used as representative of light, medium‐sized and low motile prey was occasionally consumed. Each guild could include efficient predators against pests according to its hunting strategies and the ecological exploited niches. Orb‐weavers could be efficient predators against flying pests; ambushers such as S. globosum could contribute to the reduction of the populations of flower‐visiting pests, whereas active ground hunters may also play an important role preying on pests that develop a part or all of its life cycle in the ground. However, further research on feeding behaviour such as prey switching is needed for a better understanding of the effectiveness of spiders as natural enemies.